ARTICLE longimana (Risso, 1816), a junior synonym of P. bluteli (Risso, 1816) (Crustacea: : Anomura: Porcellanidae) and a distinct from P. longicornis (Linnaeus, 1767)

Luciane Augusto de Azevedo Ferreira¹² & Marcos Tavares¹³

¹ Universidade de São Paulo (USP), Museu de Zoologia (MZUSP). São Paulo, SP, Brasil. ² ORCID: http://orcid.org/0000-0003-1683-9962. E‑mail: [email protected] ³ ORCID: http://orcid.org/0000-0002-7186-5787. E‑mail: [email protected]

Abstract. Pisidia longimana (Risso, 1816) and P. bluteli (Risso, 1816), both described from Nice, France, have been considered each other’s synonyms or have been validated depending on successive taxonomic opinions. The validity of both in respect to P. longicornis (Linnaeus, 1767) has also been contradicted a number of times. The current lack of clarity in the use of the names P. longicornis, P. longimana and P. bluteli has resulted in nomenclatural instability, but also in unreliability and miscommunication as regards the available ecological and distributional information. The validity of P. bluteli and P. longimana is revisited herein based on a large number of specimens (241 males, 190 females and 33 juveniles) from many different localities. The latter species is confirmed as a junior synonym of the former, whereas P. bluteli and P. longicornis are herein considered separate species. Diagnostic characters and morphological variations are discussed and illustrated.

Key-Words. Biodiversity; Eastern Atlantic; Mediterranean; Porcelain crabs.

INTRODUCTION tion of the available ecological and distributional information. The worldwide genus Pisidia Leach, 1820, cur- An ongoing phylogenetic analysis of the genus rently consists of 15 species distributed across Pisidia and the examination of 241 males, 190 fe- the tropical and subtropical zones of the eastern males and 33 juveniles from many different local- Pacific, western and eastern Atlantic and Indo- ities in the collections of the National Museum of West Pacific Oceans, and Mediterranean and Black Natural History, Smithsonian Institution (USNM) Seas (Haig, 1978; Osawa & McLaughlin, 2010; and Museu de Zoologia, Universidade de São Paulo Dong & Li, 2014; WoRMS, 2020). (MZUSP), prompted us to revisit the validity of Three species are currently considered valid P. bluteli and P. longimana. The latter species is con- from the eastern Atlantic and Mediterranean and firmed as a junior synonym of the former, whereas Black Seas basins: Pisidia longicornis (Linnaeus, P. bluteli and P. longicornis are herein considered two 1767), P. bluteli (Risso, 1816) and P. longima- separate species. Diagnostic characters and mor- na (Risso, 1816) (Osawa & McLaughlin, 2010). phological variations are discussed and illustrated. However, P. longimana and P. bluteli have been lumped and split alternately with each successive taxonomic opinion, and the validity of both in MATERIAL AND METHODS respect to P. longicornis has also been contradict- ed a number of times (Zariquiey-Álvarez, 1951; Abbreviations used include: cl, carapace Holthuis, 1961; García-Raso, 1987; d’Udekem length, taken from the front to the posterior me- d’Acoz, 1995, 1999; Koukouras et al., 2002; Osawa & dian margin of the carapace; cw, carapace width, McLaughlin, 2010). Thus, the current lack of clarity taken at the level of its widest point; P1, cheliped in the use of the names P. longicornis, P. longimana (pereopod 1); P2‑P4, pereopods 2 to 4; St, station. and P. bluteli has resulted in nomenclatural insta- The ecological notes and distributional rang- bility, but also in unreliability and miscommunica- es were compiled by d’Udekem d’Acoz (1999)

Pap. Avulsos Zool., 2020; v.60: e20206036 ISSN On-Line: 1807-0205 http://doi.org/10.11606/1807-0205/2020.60.36 ISSN Printed: 0031-1049 http://www.revistas.usp.br/paz ISNI: 0000-0004-0384-1825 http://www.scielo.br/paz Edited by: William Ricardo Amâncio Santana Received: 23/04/2020 Accepted: 09/06/2020 Published: 16/07/2020 Pap. Avulsos Zool., 2020; v.60: e20206036 Ferreira, L.A.A. & Tavares, M.: Pisidia longimana junior synonym of P. bluteli 2/6 separately for P. bluteli and P. longimana and are herein (USNM 1277834); Pula, Pomer Bei, from Mytilus beds, combined. 18 males, 25 females (USNM 1277836). Montenegro: Bibliographic references in the synonymic list are Sveti Juan, in bryozoans, 35 to 40 m, 28 Aug. 1990, not exhaustive, being limited only to taxonomic works 1 male (USNM 1277830). Turkey: Woflzmir, 37°25 N, ′ including nomenclatural acts and taxonomical opinions. 26°40 E, 29 to 36.5 m, M.J. Sweeney coll., 19 Jun. 1973, ′ 2 males (USNM 150831). Tunisia: Tunis, La Marsa, pub- lic beach at Marsa Plage, St. 211C, from Posidonia roots, RESULTS AND DISCUSSION M.L. Jones coll., 21 Aug. 1973, 1 ovigerous female (USNM 1277827); La Marsa, public beach at Marsa Plage, Family Porcellanidae Haworth, 1825 2 m, R.B. Manning et al. coll., 29 Jul. 1972, 1 male, 1 female Genus Pisidia Leach, 1820 (USNM 265991); La Marsa, public beach at Marsa Plage, 0 to 2 m, from sabellarid colonies; R.B. Manning & R. Type species: Cancer longicornis Linnaeus, 1767 Brown coll., 10 Aug. 1972, 22 juveniles (USNM 1278013); Salammbo, Punic Port, R.B. Manning & L.B. Holthuis coll., 04 Oct. 1972, 9 males, 3 females (USNM 265992); Pisidia bluteli (Risso, 1816) Salammbo, south port along entrance channel to Punic (Figs. 1A‑O) Port, L.B. Holthuis & R.B. Manning coll., 06 Oct. 1972, 1 male (USNM 265997); Punic Port, 24 Apr. 1974, 1 fe- bluteli Risso, 1816: 67, pl. 1, fig. 7 [type lo- male (USNM 1277832); Salammbo, flat north of and out- cality: Nice, France]; Zariquiey-Álvarez, 1951: 131, side of North Punic Port, M.L. Jones coll., 21 Aug. 1973, figs. 1, 2A‑D. 2 males, 3 females (USNM 1278014); Salammbo, North Porcellana longimana Risso, 1816: 68 [type locality: Nice, Punic Port, in channel, St. 206D, algae wash, M.L. Jones France]. coll., 18 Aug. 1973, 2 males (USNM 1278016); Salammbo, Pisidia bluteli – Haig, 1960: 209; Holthuis, 1961: 37, near Port Punic, St. 207B, grass flats, algae wash, M.L. figs. 12a, d, 13a; Zariquiey-Álvarez, 1968: 290, 291, Jones coll., 19 Aug. 1973, 5 males, 2 females, 11 juve- figs. 94e, 103a, e, f; Manning & Števčić, 1982: 297; niles (USNM 1278019); Salammbo, grass flats adjacent García-Raso, 1987: 277, fig. 7; Števčić, 1990: 229; to northern Punic Port, St. 207B, algae wash, M.L. Jones Koçak et al., 2010: 340. coll., 19 Aug. 1973, 3 males, 1 intersex (USNM 1278020); Pisidia longimana – Holthuis, 1961: 40, figs. 12b, e, 13b; Raquad, off first public beach, 2 to 4 m, R.B. Manning et al. Zariquiey-Álvarez, 1968: 291, 292, fig. 103b; Manning coll., 21 Jan. 1973, 1 male (USNM 265999); Raquad, 12 km & Števčić, 1982: 297; García-Raso, 1987: 277, north of WW II Bunker at beach proper adjacent to mouth figs. 1, 3‑5; Koukouras et al., 2002: 451, figs. 2b‑4b; of canal extending to beach, L.B. Holthuis & R.B. Manning Koçak et al., 2010: 340. coll., 10 Oct. 1972, 1 male (USNM 265993); Korbous, Pisidia forma bluteli – d’Udekem d’Acoz, 1999: 170. isolated beach between Sidi Rais and Arn Oktor, R.B. Pisidia forma longimana – d’Udekem d’Acoz, 1999: 170. Manning et al. coll., 30 May 1973, 1 male (USNM 265998); off Cap Samart Proper, 3 to 5 m, R.B. Manning coll., Material examined: Spain: Cádiz, Bay of Cádiz, Los 14 Jan. 1973, 3 males, 3 females, 3 ovigerous females Lastres, east part of Bay of Cádiz, 29 Oct. 1974, 5 m, (USNM 265994); Sidi Bou Said, south of Yatch Basin, R.B. 1 male, 1 ovigerous female (USNM 258280); Bay of Cádiz, Manning & C. Froglia coll., 29 May 1973, 11 males, 2 fe- just northwest of harbor entrance of Cádiz, 29 Oct. 1974, males, 5 ovigerous females (USNM 265995); Sidi Bou Said, 8 to 10 m, 1 male (USNM 258282); Bay of Cádiz, east just south of Yatch Basin, R.B. Manning & L.B. Holthuis part of Bay of Cádiz near Bajo de Las Cabezuelas, sand coll., 05 Oct. 1972, 2 females (USNM 265996); Sidi Bou dune on a small cape, 30 Oct. 1974, 2.5 m, 1 male Said, in Yatch Club Harbor, R.B. Manning & C. Froglia coll., (USNM 258283). Italy: Sicily, between Mazara del Vallo 23 May 1973, 13 males, 2 females, 9 ovigerous females and Capo Guanitola, 37°34 36 N, 12°39 00 E, St. Sic 85‑1, (USNM 265798); Sidi Ali El Mekki, west edge of Lac de ′ ″ ′ ″ 0.5 m, R.B. Manning & C. Froglia coll., 1985, 1 male, 1 fe- Porta Farina, behind shore about 5 km from beach (salt male (USNM 205775); Scogliere Falconara M. (Ancona), lake), St. RBM TUN‑27, R. Brown et al. coll., 13 Aug. 1972, C. Froglia coll., 11 May 1973, 0.5 m, 2 males, 1 female, 2 males, 1 ovigerous female (USNM 1277844); St. 251B, 3 ovigerous females (USNM 205865); Triestre, Grado, R.B. Manning coll., 28 Oct. 1973, 4 males, 2 females 10 m, C. Froglia coll., 16 Jan. 1974, 5 males, 2 females (USNM 1278010); St. 17, R.B. Manning coll., 4 males, (USNM 152180); Golfo di Manfredonia, 41°25 18 N, 2 ovigerous females (USNM 1277838); St. 23A, R.B. ′ ″ 16°13 30 E, 15 m, C. Froglia et al. coll., 06 Jul. 1974, Manning coll., 7 males, 6 females, 2 ovigerous females ′ ″ 8 males, 11 ovigerous females, 1 intersex (USNM 152181). (USNM 1277839); St. ALG‑9A, R.B. Manning coll., 09 Mar. Slovenia: Portoroz, 17 m, 11 Sep. 1970, 3 males, 2 fe- 1974, 2 males, 1 ovigerous female (USNM 1278009); males, 1 ovigerous female (USNM 1278008); Portoroz, St. 23A, R.B. Manning coll., 1 female (USNM 1278012); Obala, 26 Aug. 1971, 2 males, 1 female, 5 ovigerous fe- St. 211B, R.B. Manning coll., 1 male, 2 females, 1 intersex males (USNM 1278011). Croatia: Cervar, near Porec, (USNM 1278015); St. 210, R.B. Manning coll., 4 males, in Mytilus beds, 3 males (USNM 1277826); Ibidem, 1 ovigerous female (USNM 1278017); St. 207B, R.B. 4 males, 12 females (USNM 1277824); Dalmatia, under Manning coll., 4 males (USNM 1278018); St. 211C, R.B. stones, from Mytilus beds, 9 males, 7 ovigerous females Manning coll., 21 Aug. 1973, 9 males, 8 females, 2 ovig- Ferreira, L.A.A. & Tavares, M.: Pisidia longimana junior synonym of P. bluteli Pap. Avulsos Zool., 2020; v.60: e20206036 3/6

Figure 1. (A‑O) Pisidia bluteli (Risso, 1816). (A‑E) “bluteli type”, female, cl 4.6 mm, cw 4.3 mm (USNM 1278011). (F‑J) “bluteli-longimana type”, female, cl 4.4 mm, cw 4.3 mm (USNM 1278011). (K‑O) “longimana type”, female, cl 4.8 mm, cw 4.7 mm (USNM 205865). (A, F, K) carapace, dorsal view (B, G, L) left P1, dorsal view. (C, H, M) left ischial and meral P1, ventral view; (D, I, N) right antenna, dorsal view; (E, J, O) left P2, lateral view. Scale bars: (A, B, E‑G, J‑L, O) = 1 mm; (C, D, H, I, M, N) = 0.5 mm. Pap. Avulsos Zool., 2020; v.60: e20206036 Ferreira, L.A.A. & Tavares, M.: Pisidia longimana junior synonym of P. bluteli 4/6 erous females, 1 intersex (USNM 1278021); St. 38B, R.B. Algarve, Faro, south coast of Portugal, Zariquiey-Álvarez Manning coll., 1 female (USNM 1278023); St. 39B, R.B. coll., 04 Nov. 1974, 9 to 10 m, offshore, 7 males, 1 fe- Manning coll., 25 Aug. 1972, 1 male (USNM 1278024); male, 1 ovigerous female (USNM 258279); Zariquiey- St. 176B, R.B. Manning coll., 05 Jul. 1973, 1 ovigerous Álvarez coll., 04 Nov. 1974, 16 m, 3 males, 1 ovigerous female (USNM 1278025); St. 47A, R.B. Manning coll., female (USNM 258279); Armação de Pera, south coast, 2 males, 1 female (USNM 1278026). 01 Nov. 1974, 1 male (USNM 258268).

Comparative material examined: Pisidia longicornis Habitat: Pisidia bluteli has been reported from under (Linnaeus, 1767): Wales: Caernavonshire, Abedaron Bay, rocks, hard bottoms, detritic bottoms, sabellariid colo- north of Gales, O.D. John & F.C. Fraser coll., 24 Jul. 1936, nies, calcareous algae, and Mytilus beds, and rhi- 2 males, 2 ovigerous females. (USNM 156435). England: zomes of Posidonia (d’Udekem d’Acoz, 1999 and refer- Drake’s Island, Plymouth Sound, O. Hartman coll., ences therein). 16 Aug. 1939, 2 intersex (USNM 1256039); London, Channel Islands, E. Lovett coll., 5 males (USNM 6560). Distribution: This species is known from the Atlantic Germany: Helgoland, Royal Biological Station, Austern coasts of Spain, Portugal and Morocco, and the entire bânke der Nordsee, 1890, 3 males, 1 ovigerous female Mediterranean and Black Seas, between the tidal mark (USNM 19907). France: Chausey Islands, R.B. Manning & A. down to 20 m (d’Udekem d’Acoz, 1999 and references Crosnier coll., 27 Aug. 1996, 24 males, 15 females, 11 ovig- therein). erous females, 1 intersex (USNM 283078); R.B. Manning & A. Crosnier coll., 27 Aug. 1996, 2 males, 2 females, Remarks: Pisidia bluteli and P. longimana (both as 1 ovigerous female (USNM 1256055); 48°53 N, 01°50 W, Porcellana) were briefly described from Nice, France by ′ ′ north side of islands, Manning et al. coll., 27 Aug. 1992, Risso (1816). Pisidia bluteli was crudely illustrated, where- 5 males, 3 ovigerous females (USNM 264683); north side as no illustration of P. longimana was given. of Chausey Island, Manning et al. coll., 29 Aug. 1992, Pisidia bluteli was generally regarded as a junior syn- 3 males (USNM 264683); Manning et al. coll., onym of P. longicornis (Linnaeus, 1776), until Zariquiey- 28 Aug. 1992, 5 males, 1 female (USNM 264654); Granville Álvarez (1951) provided evidence that both species were et al. coll., Aug. 1996, 2 males, 2 females (MZUSP 18730); morphologically distinct. While agreeing with Zariquiey- 2 males (MZUSP 25286). Spain: Ria de Arousa, St. 1288, Álvarez, Holthuis (1961) argued that not only was P. bluteli 04 Jul. 1963, 2 males, 2 ovigerous females (USNM 121833); valid, but so was P. longimana, and he therefore removed St. 1472, 23 Jul. 1964, 1 male (USNM 121834); St. 1472, the latter species from the synonymy with P. longicornis. 23 Jul. 1964, 1 male (USNM 121835); Isla de Arousa, 0.3 km Holthuis’ (1961) view, however, was challenged by the north-northeast of Punta Campelo, 42°34,5 N, 08°52,7 W, observations of Manning & Števčić (1982), who, without ′ ′ F. Ponte coll., 18 Jul. 1964, 18 m, 2 males (USNM 121837); further details, commented that some specimens from 42°34,7 N, 08°52,3 W, St. 1607, F. Ponte coll., 20 Jul. 1964, the Piran Gulf (northern Adriatic Sea) showed intergra- ′ ′ 1 male (USNM 121840); 1.5 km north-northeast of Punta dations between P. bluteli and P. longimana. García-Raso do Campelo, 42°35,2 N, 08°52,8 W, St. 1606, F. Ponte (1987) went farther and moved P. bluteli and P. longimana ′ ′ coll., 20 Jul. 1964, 30 to 40 m, 2 males (USNM 121839); back into the synonymy with P. longicornis. Conversely, between Lobeira de Cambados and Goldferna, F. Koukouras et al. (2002), once again considered P. bluteli Ponte coll., 30 Jul. 1964, 19 m, 1 male (USNM 121847); and P. longimana as being distinct from each other and 30 Jul. 1964, 19 m, 9 males, 1 female, 1 ovigerous female from P. longicornis. (USNM 121846); 0.2 km northeast of Jidoiro Pedregoso, Arguments in favor of splitting P. bluteli from P. longima- 42°32,6 N, 08°54,8 W, St. 1924, F. Ponte coll., 12 Aug. 1964, na are essentially those of Holthuis (1961): (1) the orbital ′ ′ 5 m, 1 female (USNM 121850); O Grove, 2.2 km south- margin shows a row of spines in P. bluteli, whereas the east of Punta Canela, 42°26,6 N, 08°27,2 W, St. 1841, J. orbital margins are usually crenulate or minutely serrate, ′ ′ Cambeira coll., 06 Aug. 1964, 55 to 65 m, 1 ovigerous never spinous in P. longimana; (2) there are several distinct female (USNM 121848); 0.2 km southeast of Benencia spines on the dorsal surface of the carapace in P. bluteli, south lighthouse, 42°35,5 W‑08°52,5 W, F. Ponte coll., whereas the carapace spines are smaller and in P. longima- ′ ′ 18 Jul. 1964, 20 to 30 m, 1 male (USNM 121836); 0.6 km na larger specimens, hardly visible; (3) the antennal ba- northeast of Punta Cabio, 42°35,0 N, 08°55,3 W, St. 1631, sis-ischium and merus have a distinct spine at the distal ′ ′ F. Ponte coll., 22 Jul. 1964, 7 m, 1 male (USNM 121844); end of the mesial margin in P. bluteli, whereas the antennal 42°35,4 N, 08°54,8 W, F. Ponte coll., 04 Aug. 1964, merus bears no spine, although a distinct spine is present ′ ′ 15 to 20 m, 1 male (USNM 121849); 42°35,4 N, 08°52,6 W, in the antennal basis-ischium in P. longimana; (4) numer- ′ ′ F. Ponte coll., 20 Jul. 1964, 30 m, 1 ovigerous female ous spinules, arranged in more or less distinct longitudi- (USNM 121838); 0.3 km southwest of Los Mezos Light, nal rows, are found on the dorsal surface of the carpus 42°30,8 N, 08°55,7 W, St. 1633 (F11), F. Ponte coll., and the palm in P. bluteli, whereas the dorsal surface of the ′ ′ 22 Jul. 1964, 12 m, 1 ovigerous female (USNM 121841); carpus and palm are smooth, although in the juveniles St. 1633 (F11), F. Ponte coll., 22 Jul. 1964, 12 m, 1 male they may be provided with a median longitudinal row (USNM 121842); 0.4 km of Los Mezos Light, 42°30,7 N, of granules or spinules, in P. longimana; (5) a row of slen- ′ 08°56,0 W, St. 1634 (F12), F. Ponte coll., 22 Jul. 1964, der spinules usually is present along the lateral margin of ′ 18 to 22 m, 2 females (USNM 121843). Portugal: the carpus in P. bluteli, whereas the lateral margin of the Ferreira, L.A.A. & Tavares, M.: Pisidia longimana junior synonym of P. bluteli Pap. Avulsos Zool., 2020; v.60: e20206036 5/6 carpus is smooth in the adults, but may be provided with more spines (rarely 1) in P. bluteli, whereas, in contrast, spinules in the juveniles in P. longimana; and (6) numer- the branchial region bears 0 to1 spines (rarely 2) spines ous strong dorsal spines are present on the merus, carpus in P. longimana. However, García-Raso (1987) opined that and propodus of the walking legs in P. bluteli, whereas the the characters used by Holthuis (1961) do not allow for carpus and merus of the walking legs do not show a row distinguishing between P. longicornis, P. longimana and of spinules, although very few short and blunt granules P. bluteli for intermediate forms in which all possible com- or spinules may sometimes be observed on the merus binations of the purportedly distinguishing characters are in P. longimana (Holthuis, 1961). Additionally, Koukouras commonly found, sometimes even in the same specimen. et al. (2002) submitted that P. bluteli and P. longimana Consequently, García-Raso (1987) concluded that P. bluteli could be further differentiated in that the branchial re- and P. longimana should be sunk into the synonymy with gion, behind the cervical groove, is provided with 2 or P. longicornis (see also d’Udekem d’Acoz, 1995, 1999).The

Figure 2. (A‑E) Pisidia longicornis (Linnaeus, 1767), female, cl 5.1, cw 5.1 (MZUSP 18730). (A) carapace, dorsal view; (B) right antenna, dorsal view; (C) left P1, dorsal view; (D) left ischial and meral P1, ventral view; (E) left P2, lateral view. Scale bars: (A, C, E) = 1 mm; (B, D) = 0.5 mm. Pap. Avulsos Zool., 2020; v.60: e20206036 Ferreira, L.A.A. & Tavares, M.: Pisidia longimana junior synonym of P. bluteli 6/6 large number of specimens examined herein from the REFERENCES collections of the USNM and MZUSP lends support to the view that P. bluteli and P. longimana are synonyms. Chace, F.A. 1956. Porcellanid crabs. Expédition Océanographique Belge dans The purportedly diagnostic characters for distinguishing les eauxs côtières africaines de l’Atlantique Sud (1948‑1949). Résultats between P. bluteli and P. longimana actually intergrade Scientifiques, 3(5): 1‑54. between specimens, even from the same locality. For in- Dong, D. & Li, X. 2014. 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Zoologische Verhandelingen, of P2, P3 and P4 (Figs. 1E, J, O). 47: 1‑67. Likewise, the characters proposed by Koukouras et al. Koçak, C.; Kirkim, F. & Katağan, T. 2010. Anomuran (Crustacea, Decapoda) (2002) clearly overlap with each other and therefore, can- fauna of Fethiye Bay (Turkey, eastern Mediterranean). Turkish Journal of not be used to distinguish among P. longicornis, P. bluteli Zoology, 34: 333‑342. and P. longimana. However, three diagnostic characters Koukouras, A.; Malvidis, M. & Noel, P.Y. 2002. The genus Pisidia Leach differentiate P. longicornis from P. bluteli: (1) P. longicornis (Decapoda, Anomura) in the Northeastern Atlantic Ocean and the (as already noticed by Holthuis, 1961) presents incon- Mediterranean Sea. Crustaceana, 75(3/4): 451‑463. spicuous or absent spinulation compared to P. bluteli, Leach, W.E. 1820. Galateadées. In: Dictionnaire des Sciences Naturelles. Paris, whose spines in the carapace, antenna and P1 are always Le Normant. v. 8, p. 49‑59 well-developed (Figs. 1A‑O; 2A‑E); (2) P. longicornis males Linnaeus, C. 1767. per regna tria… 12. ed. Reformata. present the major P1 broader and swollen, whereas in Holmiae, Laurentii Salvii. v. 1, pt. 2, p. 533‑1327. P. bluteli the P1 is long, slender and slightly flattened Manning, R.B. & Števčić, Z. 1982. Decapod fauna of the Piran Gulf. Quaderni (present study); (3) the front in P. longicornis presents a del Laboratorio di Tecnologia della Pesca, 3(2‑5): 285‑304. deep longitudinal groove in the median lobe (so that the median lobe seems to be divided into two), whereas Osawa, M. & McLAughlin, P.A. 2010. Annotated checklist of anomuran P. bluteli presents three conspicuous lobes, with the lon- decapod of the world (exclusive of the Kiwaoidea and gitudinal one shallow and poorly visible (present study) families Chirostylidae and Galatheidae of the Galatheoidea). Part II. (Figs. 1A, F, K; 2A). Porcellanidae. The Raffles Bulletin of Zoology, 23: 109‑129. Pisidia longicornis s. str. is known from the Atlantic Risso, A. 1816. Histoire Naturelle des Crustacés des Environs de Nice. Paris, coast of Europe, from south Norway to Portugal, as well Librairie Grecque-Latine-Allemande. 175p. as from the Mediterranean Sea, where it inhabits greater Števčić, Z. 1990. Check-list of the Adriatic Decapoda Crustacea. Acta Adriatica, depths, between 30 and 100 m (d’Udekem d’Acoz, 1999). 31(1/2): 183‑274. Its record from the west African coast, from Mauritania Udekem d’Acoz, C. d’. 1995. Contribution à la connaissance des Crustacés to Angola (Chace, 1956) deserves further investigation. Décapodes Helléniques II: Penaeidea, Stenopodidea, Palinuridea, Homaridea, Thalassinidea et Anomura, et note sur les Stomatopodes. Bios, Macedonia (Greece), 3: 51‑77. ACKNOWLEDGMENTS Udekem d’Acoz, C. d’. 1999. Inventaire et distribution des Crustacés Décapodes de l’Atlantique nord-oriental, de la Méditerranée et des eaux We gratefully thank Rafael Lemaitre (USNM) for grant- continentales adjacentes au nord de 25°N. Patrimoines naturels, 40: ing access to the collection under his responsibility and 1‑383. for providing working space. Thanks also extend to Karen WoRMS, 2020. World Register of Marine Species: Pisidia Leach, 1820. Available at: Reed (USNM) for her kind assistance throughout the ex- http://www.marinespecies.org/aphia.php?p=taxdetails&id=106837. ecution of this work. We are also grateful to Christopher Access in: 24/05/2020. B. Boyko (American Museum of of Natural History, Zariquiey-Álvarez, R. 1951. Decapodos espanoles IV. Sobre el genero New York) for helpful suggestions. LAAF thanks CNPq Porcellana Lamarck, 1801. Publicaciones del Instituto de Biologia Aplicada, (372489/2019‑2). MT thanks CNPq (303122/2016‑1) 9: 131‑139. for supporting studies on the systematics of decapod Zariquiey-Álvarez, R. 1968. Crustáceos decápodos Ibéricos. Investigación crustaceans. Pesquera, 32: 1‑510.

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